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Zoological History
Myth, Religion and Science
The earliest antlions described in literature—scientific or otherwise—were
not the insects that we know today, but rather mythical creatures and misnamed
animals born of imaginative translation errors. Described originally in ancient
Greek texts, and later in medieval Christian bestiaries, these fictitious "ant-lions" became
the focus of fantastic stories which fulfilled symbolic or religious functions
in their respective cultures. (The convoluted history of these myths is explored
in greater detail in our section Bestiary: Creatures
of Myth and Psyche.) In the period ranging from the 11th century to the
13th, a scientific revival occurred—drawing largely upon the much earlier
work of Aristotle (384-322 B.C.E.), considered by many to be the father of
zoology. From that point on, there was a renewal in interest in scientific
research and a return to more objective methods of observation (Beckhöfer-Fialho
1996).
The path to an accurate, scientific understanding of true antlions, however, is just as fascinating—and nearly as confusing—as the history of mythical antlions. Early naturalists had to overcome translation errors and widely-varying classification attempts. Even the development of a more universally-accepted taxonomic nomenclature didn't eliminate disagreements over some antlion genus and species names—disagreements which persist up to the present day.
Ants or Antlions?
Early European scholars, typically members of the clergy, actually considered antlions to be larger species of ants (called murmêkoleontês in Greek). Aldhelm (7th century) and Hrabanus Maurus (early 9th century) both left the question open as to what sort of creatures were involved. Maurus wrote: "The Ant-Lion is a small creature, extremely hostile to ants. It conceals itself in dust and kills ants that carry provisions. It is justly called formicaleon: towards ant it behaves like a lion, although towards other creatures only like an ant" (Klausnitzer 1987).
10th-century Byzantine illustrated manuscripts, doubtless copies of much earlier documents, show an arthropod called murmêkion (Figure 1). The name indicates that the creature had something in common with ants, and may in fact portray an antlion larva (Kevan 1992).
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Figure 1. The murmêkion in a 10th-century Byzantine manuscript (Pierpont-Morgan Collection, New York). Note the clawed, jaw-like anterior appendages and the assymetrical (5 right, 4 left) legs (Kevan 1992). |
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One can say no more about these illustrations than that there may have been some early Byzantine knowledge of antlion larvae before the 10th century (Kevan 1992).
Thomas of Cantimpré, a 13th-century Dominican friar, was "one of the leading representives of anti-scientific darkness, although he studied for four years under Albertus Magnus in Cologne. But at the same time, he was a nature lover" (Klausnitzer 1987). On the subject of antlions, he writes:
This worm is of the family of Ants, but considerably larger. As long as the Ant Lion is small, it is peaceable and keeps its fury to itself. But when it becomes powerful and strong, it scorns its erstwhile associates and turns to greater ones. Once it has grown to full size and power, it lurks concealed close to the track made by ants, and lies in wait like a true brigand. When the ants go to work and return carrying some object, the Ant Lion takes it from them, strangles the ants and consumes them. In winter, it steals the food stocks that the ants have gathered in the summer, because it has procured and collected nothing for itself. Not unlike this worm are those idle persons who take from the workers what they have earned by the sweat of their brow (Klausnitzer 1987).
In the mid-13th century, Barholomaeus Anglicus referred to ant-lions in his De Proprietatibus Rerum (ca. 1240). He described an animal like a big spider, in contrast to the smaller ant: "There is another kind of spider by name "mirmicoleon" or "mirmiceon," which is also called by the name "formicaleon." It is like an ant with a white head, and it has a black body marked with white spots. And the bite of this creature is as painful as wasps. And it is called ant-lion because it hunts ants like a lion and sucks out the juices from their bodies, but is devoured by sparrows and other birds just as an ant." (Kevan 1992 and Druce 1923, pp. 350-351). An illustration of this creature was created for a 15th century bestiary (Figure 2).
Figure 2. Ant-lion and ant from a 15th-century bestiary (Cambridge University Library MS Gg 6-5), based on an account of Bartholomaeus Anglicus' mid 13th-century De Proprietatibus Rerum. (Kevan 1992 and Druce 1923).
In naming this creature, it is possible that Bartholomaeus confused "mirmicoleon" with "mirmeceon"—the
biting, spider-like myrmecion of Plinius Secundus (Naturalis Historia, Book
XXIX, C.E. 77). The latter was presumably derived from the myrmêkeiôn of
Nikandros (Kevan 1992).
True Antlions
The first unequivocal description of a true antlion larva came from Albertus Magnus in his Opus Naturarum of 1255-1270. He described the larva somewhat more accurately than his pupil: "It is not an ant as some maintain. For I have frequently observed and often pointed out to friends that this creature is similar in form to a tick. It conceals itself in sand, digging a hemispherical cavity, one pole of which is its mouth. When ants pass by gathering food, it catches and devours them. I have observed this repeatedly. In winter, they are said to plunder the food stocks of ants, because in summer, they themselves do not lay in stores of food" (Klausnitzer 1987).
The first unmistakable (if not entirely accurate) drawing comes from the Codex
Animalium of Petrus Candidus Decembrus, ca. C.E. 1460 (Figure 3) (Kevan
1992).
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Figure 3. "Ant-lion" (Formica leo),
left, and "Antlion" (Formica indica), right, from the Codex Animalium of
Petrus Candidus Decembrus, ca. C.E. 1460. The large ant is attacking a much
smaller one, either to devour or enslave it; it is a "lion" among ants. It
is not implied that the "Formica indica", an undoubted (if quadruped)
myrmeleonid larva, was actually from India; it is simply illustrative of
the fabulous creature so named (Kevan 1992). |
Linnaeus, Before and After
Antonio Vallisneri's observations on antlions were published after his death in 1730 (Kevan 1992). His illustrations (Figure 4) are not only more accurate but also very thorough, showing the various stages of an antlion's life cycle.
Figure 4. Illustrations from Antonio Vallisneri's Opere Fisico-Mediche (1733) depict life stages and pit-trap of the "Verme Formicario" (Myrmecoleon) (Kevan 1992).
Between 1738 and 1742 Réaumur published several volumes of Mémoires pour servir l'Histoire des Insectes, which included an excellent series of illustrations covering the whole life history of "Formica-leo" (generally equated with Myrmeleo formicarius Linnaeus) (Kevan 1992). Two of his larvae illustrations are shown in Figure 5.
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Figure 5. Larvae of the "Formica-leo" (Myrmeleon); illustration by Réaumur (1738, Mémoires pour servir l'Histoire des Insectes vol. 4. [left, young; right, older larva]). |
Swedish botonist Carolus Linnaeus (Karl von Linné), the founder of the nomenclature system that forms the basis of modern taxonomy, first mentions antlions in the second and fourth editions of his Systema Naturae (1740 and 1744). He called the insects Formicaleo, listing them under Hemerobius (which he used then and later for what are presently known as Neuroptera/Planipennia). In the 1744 edition he introduced the group name Neuroptera for the first time (Kevan 1992).
Another Swede, Hasselquist (1722-1752), visited Egypt on a study tour commissioned by Linnaeus. Describing a non-pit-digging antlion species, he wrote this in his diary:
At the pyramids. What pleased me most here was the Ant Lion which has its own kingdom in this place. They leapt about in the sand in hundreds, just like ants. They all held pebbles, sand, rough fragments of wood or some other thing between their neat pincers, with which they hurried to the homes they had built for themselves in the sand. Round about, I found many habitations made by this worm. They were thrown up in the sand like small mole-hills, about as large as two fists, and rather depressed on the top. In the centre of this area was a small hole, the size of a reed-stalk, through which they went in and out. I assailed them in their entrenchments, destroying a few of the latter in order to examine their internal structure and organization. But I deceived myself, for I did no more than destroy their defences, and they had prepared a secret path so skilfully that it was useless to seek out the inner part of the structure (Klausnitzer 1987).
August Johann Rösel von Rosenhof also described true antlions ("Ameis-Rauber") at length in his Die Monatlich-herauskommenden Insekten-Belustigung (1755) (Kevan 1992). In his paper entitled "The cunning and skilful Ant-Robber which transforms itself into a Terrestrial and Nocturnal Dragonfly or into a Terrestrial and Nocturnal Nymph and its Truly Amazing Characteristics," Rosenhof wrote:
As I have already indicated, the Ant Robber frequently lies hidden during the day at the side of the base of the pit; at night it is usually to be found in the middle, lying in wait with pincer-jaws at the ready, like a hunter waiting for game with his gun cocked. Does it perhaps know that insects are more likely to fall into the pit at night than during the day, or are they better able to avoid the trap set for them during the day? Both, it may be supposed, particularly since the Ant Robber usually constructs a new pit in the evening. Accordingly, I often watched very closely when it was having a meal, employing for this purpose a good magnifying glass; but I could at no time perceive any other instrument except its pincers alone, with which it seemed to suck dry its prey, holding it up out of the sand with their two extreme tips: if there had been anything else there, I could not have failed to perceive it, for these two tips protrude a long way from the head. There is then nothing apart from the two parts of the pincers themselves, which presumably must be hollow. . . Since there is no doubt that our insect draws in its food through its prehensile pincers, the question arises—for as I said above, no rectal passage exists—as to the means by which the waste part of the food is removed from the body. All insects so far described by me have a rectum, which you can quite easily see if you squeeze the abdomen slightly, when it will protrude from the hind segment. I sought it for a long time, but was unable to find it; yet neither could I imagine that the insect predators did not have to excrete waste matter, particularly since I know that spiders, for example, which also suck only the juices from their prey, are nevertheless obliged to excrete the waste portion of their food. Could it be that everything our insect ingests is used up to its benefit and growth, or is a remnant of it lost by evaporation, as Herr Réaumur believes? This certainly appears to be the case, for even that great biologist was unable to discover the Ant Lions discharge any excrement. To this end, he fed them copiously with large quantities of midges, and when they had eaten their fill and seemed replete, and indeed looked thoroughly satiated, he placed them on a clean porcelain dish, but found no indication whatever that they gave off waste matter (Klausnitzer 1987).
See Rosenhof's illustrations of eggs, pupae, larvae and adults.
In the Twelfth Edition of Systema Naturae (1767) Linnaeus first proposed the generic name Myrmeleon (Kevan 1992).
Modern Nomenclature
The history of modern antlion nomenclature is extremely complicated and is described by Kevan (1992) in more detail than can be conveyed here. In short, there persists much confusion over the use of the name formicaleo. One scholar, Leraut, attempted in 1980 to overcome the confusion by proposing that the International Commission on Zoological Nomenclature should use its powers to suppress the names Hemerobius formicaleo and H. formicalynx, both of Linnaeus, and to validate Myrmeleon formicarium(-us) of Linnaeus. As of 1991 no action had been taken regarding this proposal (Kevan 1992).
References
Beckhöfer-Fialho, Aura. 1996. "Medieval Bestiaries and the Birth of Zoology." Originally published as a website.
Druce, G. C. 1923. [Myrmekoleon] or Ant-Lion. Pp. 347-363 in The Antiquaries Journal. Vol. III, No. 4. Oxford University Press, London.
Kevan, D. K. McE. 1992. Antlion ante Linné: [Myrmekoleon] to Myrmeleon (Insecta: Neuroptera: Myrmeleonidae [sic]). Pp. 203-232 in Current Research in Neuropterology. Proceedings of the Fourth International Symposium on Neuropterology [Symposium held in Bagnères-de-Luchon, France, 1991.] M. Canard, H. Aspöck, and M. W. Mansell, eds. Toulouse.
Klausnitzer, Bernhard. 1987. Insects: Their Biology and Cultural History. New York: Universe Books.
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